The colonial cyanobacterium Trichodesmium as a physical and nutritional substrate for the harpacticoid copepod Macrosetella gracilis

The pelagic harpacticoid copepod Macrosetella gracilis usesthe colonial cyanobacterium Trichodesmium not only as a physicalsubstrate for juvenile development, but also as a food source.By associating itself with a buoyant colonial cyanobacterium,M.gracilis has developed a successful mode of life for existencein the plankton. Further evidence of M.gracills' dependenceon Trichodesmium as a physical substrate is demonstrated bypreviously undescribed microscopic observations of a gravidM.gracilis female attaching eggs to a Trichodesmium colony.Shipboard experiments investigating the ingestion and assimilationof Trichodesmium carbon (C) were conducted in September 1991and January/February 1992 in waters of the Bahamas and the Caribbean,respectively. Macrosetella gracilis not only ingested, but rapidlyincorporated, cyanobacterial organic matter into its own cellularmaterial. Utilization of ingested Trichodesmium by M.graciliswas investigated by assessing the metabolic partitioning andincorporation of ¹⁴C-labelled Trichodesmium into copepod lipids,proteins, polysaccharides and low-molecular-weight (LMW) compoundsusing sequential biochemical fractionation techniques. Despitevariations in grazing rates between the two sites and times(September 1991,0.017 µg C^* µg^–1 C h^–1;January 1992, 0.134 µg C ^* µg^–1 C h^–1,the partitioning of incorporated C into the different biochemicalfractions was relatively consistent. There was rapid assimilationof ingested C into the LMW ({small tilde}60%) and polysaccharidefractions ({small tilde}30%) in the first few hours, with asubsequent increase in the percent C incorporated into protein.On average, {small tilde}21% of the Trichodesmium C ingestedby M.gracilis was assimilated. Therefore, M.gracilis is an importantsecondary link in the food web of oligotrophic waters whereTrichodesmium is abundant.     

Localization and Characterization of a Novel 20 kDa Polypeptide in the Chloroplast of the Green Alga Dunaliella salina

Recent work with the green alga Dunaliella salina showed thepresence of a {small tilde}20 kDa chloroplast protein that wasrecognized by polyclonal antibodies raised against the isolatedLHC-II [Webb M.R. and Melis A. (1995) Plant Physiol. 107: 885].In this report, a characterization of the {small tilde}20 kDapolypeptide is presented. It is shown that it is localized inthe chloroplast envelope membrane of D. salina. The abundanceof this protein is constant on a per cell basis and independentof the light regime during cell growth. The {small tilde}20kDa polypeptide is easily degraded to a {small tilde}19 kDaproduct during sample preparation. A limited amino acid sequenceof 21 residues from the free N-terminus of the {small tilde}19kDa product was obtained. On the basis of this partial sequence,it was concluded that the {small tilde}20 kDa polypeptide isnot a degradation product of a known LHC-II but rather a novelprotein. The {small tilde}20kDa polypeptide did not cross-reactwith antibodies raised against the Cbr (carotene biosynthesis-related)gene product and showed a different electrophoretic mobilityfrom the latter. Light-shift experiments suggest that the {smalltilde}20 kDa polypeptide is not an ELIP (early light-inducibleprotein). Possible functions of the {small tilde}20 kDa proteinare discussed. ¹Permanent address: Department of Biochemistry, University oflund, PO Box 124, S-221 00 Lund, Sweden     

Diurnal vertical migration of Gymnodinium mikimotoi during a red tide in Hoketsu Bay, Japan

The depth, velocity and timing of the vertical migration ofGymnodinium mikimotoi were examined using fluorometric fieldobservations. Gymnodinium mikimotoi can migrate vertically {smalltilde}20 m daily at velocities of {small tilde}2.2 m h^–1     

Nitrate and ammonium uptake by plankton in an Amazon River floodplain lake

Uptake of ammonium and nitrate by plankton was measured in tropicalLake Calado, Brazil. Nitrate uptake was strongly influencedby light and was light saturated at {small tilde}340 µEm^–2 s^–1. In contrast, uptake of ammonium was lessinfluenced by light, and saturated at {small tilde}250 µEm^–2 s^–1. Uptake rates of both forms of nitrogenwere inhibited by up to 80% at light intensities higher thanthose required for saturation. Concentrations of ammonium andnitrate also had a strong influence on uptake rates. Half-saturationconstants (0.3–5 µM) were usually greater than ambientconcentrations (0.1–0.6 µM), indicating that uptakerates at ambient concentrations were less than one-half of thesaturated rates. Ammonium is the more important type of inorganicnitrogen for plankton of Lake Calado because nitrate concentrationsremain low to undetectable except during periodic inputs ofnitrate-rich water from the Amazon River. Using the observeddependence of uptake on concentration and light, maximum uptakerates per unit chlorophyll were computed to be in reasonableagreement with rates derived from P^B_m values for carbon uptake. ¹ Present address: Florida Department of Natural Resources,Marine Research Laboratory, St Petersburg, FL 33701, USA     

The annual cycle of Mesodinium rubrum in the waters surrounding the Isles of Shoals, Gulf of Maine

Mesodinium rubrum was collected in Kemmerer bottles, fixed inBouin's solution and protargol stained. Cell volume showed aseasonal change of over an order of magnitude, being largest(3.25x10⁴ µm³) in the early spring and smallest in thesummer. Cell abundance was highest in the spring and lowestin the summer. Biomass followed a similar trend ranging from{small tilde}1 to 147 J m^–3 Production, estimated by amultiple regression that incorporated ambient temperature andcell volume, was 2.5 kJ m^–3 day^–1 This is {smalltilde}0.3% of the primary production.     

The initiation of Phaeocystis colonies

This study was designed to elucidate the sequence of eventsthat leads to the formation of new colonies of Phaeocystis sp.(strain PCC 540) starting from single cells released from maturecolonies. Colonies were first isolated by filtration onto a10 µm mesh. Colonial cells were then liberated by shakingand inoculated into individual culture wells containing mediumwith a PO₄^2– concentration of {small tilde}1 µM.Cell size and shape were determined daily by image analysis,while chlorophyll and DNA distributions were estimated by flowcytometry. Released cells were non-flagellated and mostly locatedin the G₁ phase of the cell cycle. They developed flagella andup to 90% became motile within 24 h. Swarmers lost motilityrapidly, became elongated, began to cycle again, excreted amucilaginous compound and divided leading to new colonies withina few days. During this reproducible process, no change of ploidycould be observed. Colonies initially adhered to the bottomof culture wells. Frequent mixing drastically reduced the fractionof colonies produced and their volume. High initial PO₄^2–concentrations (5 µM) delayed colony appearance, whereaslow concentrations (0.3 µM) prevented colony formation.The two main conclusions of this study are: (i) under favorableconditions ({small tilde}1 µM PO₄^2– no mixing),a large percentage of released colonial cells give back coloniesafter going through a flagellated stage; (ii) sexuality doesnot appear to be involved in this process. ¹Present address: CREMA BP 5, F-17137 L'Houmeau, France     

Utilization of limiting concentrations of ortho-phosphate and production of extracellular organic phosphates in cultures of marine diatoms

The utilization of ortho-phosphate by two coastal marine diatomspecies, Nitzschia closterium and Cyclotella cryptica, was studiedin batch cultures. The hypothesis was tested that thresholdconcentrations in the phosphate uptake determine the lower limitof environmental phosphate, permitting the existence of species.The turn-over time of residual medium phosphate in culturesis {small tilde}10 min, indicating a rapid equilibration ofconcentration dependent on uptake with leakage of ortho-phosphate.Increasing phosphate starvation in cultures diminished the residualortho-phosphate in the range of {small tilde}60–<2nmol l^–1, as measured radiochemically after elution onSephadex® G-10 gel. These concentrations encompass the rangeof limiting phosphate concentration in continuous cultures ofthe few microalgae, for which these concentrations are actuallymeasured. The diatoms excreted {small tilde}20–100 nmolI^–1 of organic phosphate. One dominating compound, probablyan unusual nucleotide, was incompletely or not resorbed underphosphate starvation. In contrast, Nitzschia closterium excretedunder ample phosphate supply a series of three related compounds,probably phospholipids, that were resorbed under depletion.The association of the organic phosphates with macromolecularexudates is interpreted, along with the other observations,as an indication for a hardly explored periplasmatic phosphatemetabolism in these algae. ³Dedicated to Prof. Dr. H.-A. von Stosch in honour of his 75thbirthday. ⁴This study was conducted at the University of Marburg undersupport of the Humboldt Foundation Publication no. 64 of theproject "Biological Research of the Eems-Dollard Estuary".     

Relation between Ultra-Violet Spectral Change and Nucleotide Binding of the Chloroplast Coupling Factor 1

Properties of the nucleotide binding sites on chloroplast couplingfactor 1 (CF₁) were studied by equilibrium dialysis and UV spectroscopy.From our direct binding studies, we identified at least fourkinds of ADP binding sites on CF₁; a barely dissociable ADPbinding site (site A), a slowly exchangeable high affinity sitewith dissociation constant (Kd) 0.021 µM (site B), anotherslowly exchangeable high affinity site with Kd 1.6 µM(site C) and several low affinity (Kd {small tilde}30 µM)sites. The Kd values for sites B and C of the other nucleotidestested were 0.5 µM and 16 µM (GDP), 8 µM and34 µM (CDP), 17 µM and 20 µM (UDP) and 1.4µM and 1.4 µM (PP₁). From a comparison of the observed UV spectral change and theamount of nucleotide bound to these sites, as calculated fromthe above Kd values, we concluded that the nucleotide bindingto site B or G induces UV spectral changes that are almost thesame in shape and magnitude. The estimated difference molarabsorption coefficient () was 3.4?10³M^–1_ADP cm^–1for ADP at 278 nm. Our conclusions were strengthened by thegood agreement between the observed spectra and the calculatedspectra (derived from the Kd and values of ADP and GDP) whenADP and GDP were added together to CF₁. The cause of the unusual behavior of GDP in the UV differencespectrum which was unexplained in our previous report was shownto be competition between the GDP added and previously boundADP at sites B and C; this distorted the real spectrum inducedby GDP. (Received October 3, 1983; Accepted February 13, 1984)     

Annual pattern of micro- and mesozooplankton abundance and biomass in a subtropical estuary

The pattern of biomass and abundance of microzooplankton andmesozooplankton were studied over an annual cycle in the NuecesEstuary, Texas. Zooplankton samples and associated hydrographicdata were collected at four locations at biweekly intervalsfrom September 1987 through October 1988. This is a broad, shallowbay system with an average depth of 2.4 m. The concentrationof chlorophyll a in the surface waters averaged 7.4 µgl^–1with 85% passing through a 20 µ mesh. Microzooplankton(20–200 µ in length) were extremely abundant throughoutthis study. Abundances of ciliates (including both aloricateciliates and tintinnids) ranged from 5000 to 400 000 l^–,with a mean of 38 000 l^–1 of seawater over the entirecourse of the study. Mesozooplankton (200–2000 µmin length) abundance averaged 6100 m^–3 for samples collectedduring the day and 10 100 m^–3 for samples collected atnight. Mesozooplankton were dominated by Acartia tonsa whichmade up {small tilde}50% of the total. Biomass estimates formicrozooplankton (based on volume estimates) were often higherthan measured biomass of mesozooplankton. Given the shortergeneration times and higher metabolic rate of microzooplanktoncompared to mesozooplankton, microzooplankton should have agreater effect on the trophic dynamics of the Nueces Estuarythan mesozooplankton.     

Energetics and feeding dynamics of Euphausia superba in the South Georgia region during the summer of 1994

Measurements of adult Antarctic krill (Euphausia superba) gutcontents, evacuation and egestion rates, as well as digestiveefficiency, were carried out during February-March 1994 in thevicin ity of South Georgia to estimate in situ daily ration.These were combined with acoustically derived biomass data tocalculate the grazing impact of Antarctic krill and its contributionto the carbon flux in the region. Individual levels of gut pigmentconcentrations and evacuation rates ranged from 27 to 1831 ngchlorophyll a-eq. ind.^–1 and from 0.133 to 0.424 h^–1,respectively. Losses of pigment fluor escence during digestionwere very high, ranging from 58 to 98% of the total pigmentdigested. Daily carbon consumption estimated using the gut fluorescencemethod varied from 0.234 to 0.931 mg C ind.^–1 day^–1(or 0.4–1.7% of body carbon), compared to {small tilde}2.73mg C ind.^–1 day^–1 (or {small tilde}5% of body carbon)using the faecal pellet production data. The 3-fold higher dailyration estimated using egestion rate data may be explained bypredation on micro-and mesozooplankton. Maximum krill grazingimpact ranged from 0.4 to 1.9% of the total phytoplankton stockor from 10 to 59% of the total daily primary production. However,grazing impact on the microphytoplankton (>20 µm) wassubstantially higher, at times exceeding 100% of the daily microphytoplanktonproduction. It is suggested that to meet its energetic demands,kriil must consume a substantial proportion of heterotrophiccarbon. ³Present address Zoology Department, University of Fort Hare,P/Bag X1314, Alice, 5700, South Africa     

Summer nutrient dynamics in the Middle Atlantic Bight: primary production and utilization of phytoplankton carbon

In late summer, production and utilization of carbon in thestratified water of the Middle Atlantic Bight appears to approachsteady-state conditions. In the euphotic zone there is a paniculateorganic carbon (POC) pool of {small tilde}6000 mg Cm^–2.Primary production adds {small tilde}350 mg C m^–2 d^–1,while zooplankton ingestion removes {small tilde}450 mg C m^–2d^–1, of which {small tilde}135 mg C m^–2 d^–1are returned to the POC pool as feces. Sinking of POC averages{small tilde}240 mg C m ^–2 d^–1. Thus, there is anet loss from the euphotic zone of {small tilde}200 mg C m^–2d^–1, which represents a removal rate from the POC poolof {small tilde}3%d^–1. However, sinking losses of phytoplanktoncarbon from the POC pool were small ({small tilde}12 mg C m^–2d^–1), which suggests that most of the primary productionenters the pelagic food chain. This is in sharp contrast tothe conditions in early spring, when the POC pool increasessubstantially, and the main loss is sinking rather than heterotrophicconsumption. ^*This research was supported by the U.S. Department of Energyunder Contract No. DE-AC02-76CH000I6.     

Laboratory study of the spawning rate of the calanoid copepod Centropages typicus: effect of fluctuating food concentration

The spawning rate of laboratory-reared Centropages typicus fedHymenomonas elongala increases with food concentration, up toa value of {small tilde}2800 µg C (16 500 cells) ml^–1.An alternation of a low food (1000 cells ml^–1) and highfood concentration (16 500 cells ml^–1) is not favourableto egg release when its periodicity is 1 or 2 days, whereasit may be of advantage if it is longer (3–6 days). Inthe latter case, Centropages typicus will benefit best fromthe rich food diet if this coincides with (or just follows)the last moult.     

Studies on the Movement of Water Through Apple Trees

Resistances to the flow of water through young potted appletrees were estimated by measuring the transpiration rate oftrees with and without root systems. Root system resistanceswere obtained by difference. Whole-plant resistances were ofthe order 10 x 10¹³ Pa s m^–3 and there was some evidencethat root resistances (R_r) varied with transpiration rate; theratio R_r:R_x (where R_x is resistance to water flow in the stemsystem) altered from 2:1 at relatively high transpiration ratesto 1:1 at lower rates. The trunk of a 9-year-old orchard tree (trunk diameter {smalltilde}7 cm, height {small tilde}2.5 m) was cut under water andestimates of the flow resistances in this tree were obtained.These were much lower than the resistances to flow in the pottedtrees. Capacitance (defined as the change in stored water content perunit change in plant water potential) values were calculatedfor the small trees and the large tree from measurements ofweight and water potential changes after the trees were removedfrom water. They were very similar on a weight basis (approx.2.0 x 10^–8 kg kg^–1 Pa^–1). Leaf capacitancevalues ({small tilde}1 x 10^–8 kg Pa^–1 m^–2)were also obtained. Stomatal conductances decreased with water potential and increasedwith short-wave radiation, but the relationships were not definitive.Estimates of boundary layer conductance in a greenhouse (verylow wind speeds) were of the same order ({small tilde}5 mm s^–1)as values obtained previously.     

Specific growth rates of heterotrophic plankton organisms in a eutrophic lake during a spring bloom

The in situ growth of the dominating pelagic organisms at severaltrophic levels was investigated during a spring bloom characterizedby well-mixed cold water. The study includes primary productionand the carbon flow through the nano-, micro- and mesozooplanktonpopulations based on population dynamics and specific growthrates. The phytoplankton biomass and production were totallydominated by small algae <20 µm. of which {small tilde}5%were <3µm. potentially a food source for the nano-and microzooplankton. The mean carbon-specific primary productionwas 0.15 day^–1 and was regulated solely by light. Themean volume-based specific growth rate of bacterioplankton wasmodest. 0.1 day^–1. and probably controlled by the lowtemperature. The volume-based specific growth rates of heterotrophicnanoflagellates. ciliates. rotifers and copepods were 0.35.0.13. 0.16 and 0.03 day^–1, respectively. The observedgrowth of the heterotrophic plankton was generally not foodlimited, but was controlled by temperature. The stable temperatureduring the experiment therefore allows a cross-taxonomic comparisonof specific growth rates. The b exponent in the allometric relationship(G = aV^th) between volume-specific growth rate (G) and individualbody size (V) was –0.15 ± 0.03 for all filtratingzooplankton. indicating an in situ scaling not far from thephysiological principles onginally demonstrated for laboratorypopulations.     

Autotrophic picoplankton in southern Lake Baikal: abundance, growth and grazing mortality during summer

Autotrophic picoplankton were highly abundant during the thermalstratification period in late July in the pelagic area (waterdepth 500–1300 m) of southern Lake Baikal; maximum numberswere 2 x 10⁶ cells ml^–1 in the euphotic zone ({small tilde}15m). Unicellular cyanobacteria generally dominated the picoplanktoncommunity, although unidentified picoplankton that fluorescedred under blue excitation were also abundant (maximum numbers4 x 10⁵ cells ml^–1) and contributed up to {small tilde}40%of the total autotrophic picoplankton on occasions. Carbon andnitrogen biomasses of autotrophic picoplankton estimated byconversion from biovolumes were 14–84 µg C l^–1and 3.6–21 µg N l^–1. These were comparableto or exceeded the biomass of heterotrophic bacteria. Autotropicpicoplankton and bacteria accounted for as much as 33% of paniculateorganic carbon and 81% of nitrogen in the euphotic zone. Measurementsof the photosynthetic uptake of [^l4C]bicarbonate and the growthof picoplankton in diluted or size-fractionated waters revealedthat 80% of total primary production was due to picoplankton,and that much of this production was consumed by grazers inthe <20 µ.m cell-size category. These results suggestthat picoplankton-protozoan trophic coupling is important inthe pelagic food web and biogeochemical cycling of Lake Baikalduring summer.     

Short-term changes of protozoan control on autotrophic picoplankton in an oligo-mesotrophic lake

In May 1994, we investigated the short-term development of theplanktonic community in the epi- and metalimnion of an oligo-mesotrophiclake (Piburger See, Tyrol), focusing on trophic links betweenprotists and picoplankton, but also including phyto- and zooplankton.Uptake experiments with fluorescently labelled bacteria (FLB)and picocyanobacteria (FIC) were performed in order to comparethe importance of both prey types as carbon sources for bacterivorousprotists. Heterotrophic nanoflagellates (HNF) were responsiblefor {small tilde}90% of total protozoan picoplanktivory (FLB+ FLC); ciliates accounted for {small tilde}10%. A selectivityindex related to prey density showed that both HNF and ciliatesclearly preferred FLC over FLB. The mean cell size of autotrophic(prokaryotic) picoplankton (APP) was nearly three times larger(0.323 µm³) and much less variable than mean bacterialcell volume (0.122 µm³). Although APP biomass was on averageonly 8.6% of total picoplankton biomass, pico-cyanobacteriaaccounted for a mean 15.9% of total HNF carbon uptake. We calculatedthat total HNF grazing could match potential APP maximum growthrates at the beginning of the study period. A strong decreasein HNF individual clearance rate (CR) on APP was clearly relatedto a fall in the percentage of choanofiagellates (from 75 to{small tilde}10% of the HNF community). A simultaneous decreasein HNF biomass and CR was followed by a steep increase in APPabundance; APP abundance and HNF biomass were highly negativelycorrelated both in the epi- and the metalimnion (r_{1 EM} = –0.879,r_{1 META} = =0.907; P = 0.001). Total rotifer abundance increasedby a factor of 50 within 2 weeks and was also negatively correlatedwith HNF biomass (r_{1 EM} = –0.852, P < 0.001; r_{1 META}= –0.659, P < 0.05). HNF grazing was found to exerta strong short-term control on picocyanobacteria and this linkwas probably broken by an increase in metazooplankton, especiallydue to rotifer predation on HNF.     

Abundance and diversity of surface zooplankton in the Gulf of Aqaba, Red Sea, Egypt

Surface zooplankton were studied in Egyptian coastal watersof the Gulf of Aqaba, from bimonthly samples from July 1994to May 1995. Species diversity, numerical abundance and dynamicswere analysed for each taxon, at six sites, inside three Protectorates.A total of 62 taxa and species were identified. At all sites,copepods were predominant in the standing crop with an averageof 1945 ind. M^–3 and formed {small tilde}75.5%, numerically,of the total zooplankton community. The meroplanktonic larvaeoccupied the second rank and they constituted {small tilde}19.7%of the total zooplankton. Seasonally, the main peak of zooplanktonabundance was recorded in winter (January) with an average of3510 ind. M^–3 while September was characterized by thelowest density (1906 ind. m^–3 The relatively higher diversityvalues were recorded at Ras Mohammed Protectorate and a progressivedecline in diversity was observed northward.     

Diel variations in gut pigment content, diel vertical migration and estimates of grazing impact for copepods in the southern Benguela upwelling region in October 1987

The gut fluorescence technique was used to estimate ingestionand filtration rates of the adult female copepods Paracalanusparvus, Cenlropages brachiatus and Calanus austrails, and copepoditestages 3, 4 and 5 of C.australis in the southern Benguela upwellingregion. During the study period chlorophyll concentrations withinthe upper 20 m of the water column were high, 5 µg I^–1in mid-shelf waters and 15–30 µg I^–1 in innershelf waters. Copepod gut pigment content was low and constantduring the day then increased sharply during the first 2 h aftersunset. Gut pigment content was 2–6 times higher duringthe night compared with daytime values. Small non-migratingcopepods (Paracalanus parvus) showed the smallest diel differencein gut pigment content and large migrating copepods (Centropagesbrachiatus and Calanus australis) the largest difference. Eggproduction rates were 20 and 50% of maximum at the mid-shelfand inner shelf stations respectively, suggesting food-limitation.Comparison of ingestion rates calculated from egg productiondata with ingestion rates calculated from gut pigment data suggestedthat the copepods were feeding omnivorously at the inner shelfstations but herbivorously at the mid-shelf stations. Assumingthat all of the phytoplankton was available as food, the nearshorecopepod assemblage grazed {small tilde}1% of the standing cropeach day, and the mid-shelf assemblage grazed 5% day^–1.Because of errors and uncertainties associated with the gutfluorescence technique, the feeding impact could be underestimatedby 2–4-fold. We discuss several approaches which couldlead to more precise estimates of feeding rates. ³Present address: Marine Sciences, SUNY, Stony Brook, NY, 11794-5000,USA     

Photosynthetic characteristics of Dinophysis norvegica Claparede & Lachmann, a red-tide dinoflagellate

The relationships between photosynthesis and photosyntheticphoton flux densities (PPFD, P-l) were studied during a red-tideof Dinophysis norvegica (July-August 1990) in Bedford Basin.Dinophysis norvegica, together with other dinoflagellates suchas Gonyaulax digitate, Ceratium tripos, contributed {small tilde}50%of the phytoplankton biomass that attained a maximum of 16.7µg Chla 1^– and 11.93 10⁶ total cells I^–1.The atomic ratios of carbon to nitrogen for D.norvegica rangedfrom 8.7 to 10.0. The photosynthetic characteristics of fractionatedphytoplankton (>30 µm) dominated by D.norvegica weresimilar to natural bloom assemblages: o (the initial slope ofthe P-l curves) ranged between 0.013 and 0.047 µg C [µgChla]^–1 h–1 [µmol m^– s^–1]^–1the maximum photosynthetic rate, p^B_m, between 0.66 and 1.85µg C [µghla]^–1 h^–1; l_k (the photoadaptationindex) from 14 to 69 µ,mol m^–2 s^–1. Carbonuptake rates of the isolated cells of D.norvegica (at 780 µmolm^–2 s^–1) ranged from 16 to 25 pg C cell^–1h^– and were lower than those for C.tripos, G.digitaleand some other dinoflagellates. The variation in carbon uptakerates of isolated cells of D.norvegica corresponded with P^B_mof the red-tide phytoplankton assemblages in the P-l experiments.Our study showed that D.norvegica, a toxigenic dinoflagellate,was the main contributor to the primary production in the bloom.